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How Does Mindfulness Meditation Work? Proposing Mechanisms of Action From a Conceptual and Neural Perspective Britta K. Hölzel, Sara W. Lazar, Tim Gard, Zev Schuman-Olivier, David R. Vago and Ulrich Ott Perspectives on Psychological Science 2011 6: 537 DOI: 10.1177/1745691611419671

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Downloaded from pps.sagepub.com at The University of Iowa Libraries on May 24, 2012 Perspectives on Psychological Science 6(6) 537 –559 How Does Mindfulness Meditation Work? © The Author(s) 2011 Reprints and permission: sagepub.com/journalsPermissions.nav Proposing Mechanisms of Action From a DOI: 10.1177/1745691611419671 Conceptual and Neural Perspective http://pps.sagepub.com

Britta K. Hölzel1,2, Sara W. Lazar2, Tim Gard1,2, Zev Schuman-Olivier2, David R. Vago3, and Ulrich Ott1 1Bender Institute of Neuroimaging, Justus Liebig-University, Giessen, Germany; 2Massachusetts General Hospital, Harvard Medical School, Boston, MA; and 3Brigham and Women’s Hospital, Harvard Medical School, Boston, MA

Abstract Cultivation of mindfulness, the nonjudgmental awareness of experiences in the present moment, produces beneficial effects on well-being and ameliorates psychiatric and stress-related symptoms. Mindfulness meditation has therefore increasingly been incorporated into psychotherapeutic interventions. Although the number of publications in the field has sharply increased over the last two decades, there is a paucity of theoretical reviews that integrate the existing literature into a comprehensive theoretical framework. In this article, we explore several components through which mindfulness meditation exerts its effects: (a) attention regulation, (b) body awareness, (c) emotion regulation (including reappraisal and exposure, extinction, and reconsolidation), and (d) change in perspective on the self. Recent empirical research, including practitioners’ self-reports and experimental data, provides evidence supporting these mechanisms. Functional and structural neuroimaging studies have begun to explore the neuroscientific processes underlying these components. Evidence suggests that mindfulness practice is associated with neuroplastic changes in the anterior cingulate cortex, insula, temporo-parietal junction, fronto-limbic network, and default mode network structures. The authors suggest that the mechanisms described here work synergistically, establishing a process of enhanced self-regulation. Differentiating between these components seems useful to guide future basic research and to specifically target areas of development in the treatment of psychological disorders.

Keywords anxiety disorders, attention, cognition, consciousness, neuroscience, positive psychology, stress disorders

Mindfulness meditation has been reported to produce beneficial treatment of disorders and improvement of health; it has also effects on a number of psychiatric, functional somatic, and been shown to produce positive effects on psychological well- stress-related symptoms and has therefore increasingly been being in healthy participants (Carmody & Baer, 2008; Chiesa & incorporated into psychotherapeutic programs (cf., Baer, 2003; Serretti, 2009) and to enhance cognitive functioning (Jha, Grossman, Niemann, Schmidt, & Walach, 2004). A large body Krompinger, & Baime, 2007; Ortner, Kilner, & Zelazo, 2007; of research documents the efficacy of mindfulness-based inter- Pagnoni & Cekic, 2007; Slagter et al., 2007). Historically, mind- ventions in the treatment of a number of clinical disorders, fulness is a concept stemming from ancient Buddhist philoso- including anxiety (Hofmann, Sawyer, Witt, & Oh, 2010; phy (Bhikkhu, 2010), and is practiced to achieve enduring Roemer, Orsillo, & Salters-Pedneault, 2008), depression (Hof- happiness (Ekman, Davidson, Ricard, & Wallace, 2005) and to mann et al., 2010; Teasdale et al., 2000), substance abuse gain insight into a view of the true nature of existence (Olen- (Bowen et al., 2006), eating disorders (Tapper et al., 2009), and dzki, 2010). chronic pain (Grossman, Tiefenthaler-Gilmer, Raysz, & Kesper, 2007). Furthermore, mindfulness meditation positively influ- ences aspects of physical health, including improved immune function (Carlson, Speca, Faris, & Patel, 2007; Davidson et al., 2003), reduced blood pressure and cortisol levels (Carlson et al., Corresponding Author: 1 Britta K. Hölzel, Bender Institute of Neuroimaging, Justus Liebig-University, 2007), and increased telomerase activity (Jacobs et al., 2010). Otto-Behaghel-Str. 10 H, 35394 Giessen, Germany Not only has mindfulness successfully been used in the E-mail: [email protected]

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Table 1. Previous Theoretical Accounts That Describe Mechanisms of Mindfulness Meditation

Publication Suggested components Shapiro, Carlson, Astin, and Freedman (2006) Attention, intention, attitude Brown, Ryan, and Creswell (2007) Insight, exposure, nonattachment, enhanced mind–body functioning, integrated functioning Baer (2003) Exposure, cognitive change, self-management, relaxation, acceptance Five Facet Mindfulness Questionnaire (Baer, Smith, Hopkins, Observing, describing, acting with awareness, nonjudging of inner Krietemeyer, & Toney, 2006) experience, nonreactivity to inner experience

Definition of Mindfulness including (a) insight, (b) exposure, (c) nonattachment, (d) enhanced mind–body functioning, and (e) integrated function- In current research contexts, mindfulness is typically defined ing. Similarly, in her 2003 review, Ruth Baer summarized sev- as nonjudgmental attention to experiences in the present eral mechanisms that may explain how mindfulness skills can moment (Kabat-Zinn, 1990). Bishop et al. (2004) suggest a lead to symptom reduction and behavior change, namely (a) two-component model of mindfulness, where the first compo- exposure, (b) cognitive change, (c) self-management, (d) nent is the regulation of attention in order to maintain it on the relaxation, and (e) acceptance. A valuable empirical account immediate experience, and the second component involves for the description of the facets of mindfulness is the Five approaching one’s experiences with an orientation of curiosity, Facet Mindfulness Questionnaire (Baer, Smith, Hopkins, Kri- openness, and acceptance, regardless of their valence and etemeyer, & Toney, 2006), which was developed based on an desirability. Mindfulness is typically cultivated in formal med- item pool of previously existing mindfulness questionnaires. itation practices, such as sitting meditation, walking medita- Factor analyses over these items yielded five facets of mind- tion, or mindful movements (Kabat-Zinn, 1990). The practice fulness: observing (attending to or noticing internal and exter- of mindfulness meditation encompasses focusing attention on nal stimuli, such as sensations, emotions, cognitions, sights, the experience of thoughts, emotions, and body sensations, sounds, and smells); describing (noting or mentally labeling simply observing them as they arise and pass away. these stimuli with words); acting with awareness (attending to one’s current actions, as opposed to behaving automatically or absentmindedly); nonjudging of inner experience (refraining Need for a Theoretical Framework from evaluation of one’s sensations, cognitions, and emo- It is striking that this seemingly simplistic practice can have tions); and nonreactivity to inner experience (allowing such a wide range of applications and effects. Along with the thoughts and feelings to come and go, without attention get- many positive implications of mindfulness arises the question: ting caught in them). How does mindfulness work; what are its mechanisms? The field has produced a number of studies utilizing psycho- Although there is currently a large body of literature, covering logical scales or behavioral tasks that provide empirical support a wide range of research, including qualitative research, feasi- for some of the proposed components. Simultaneously, a grow- bility trials, controlled clinical trials, behavioral studies, and ing body of neuroimaging literature begins to describe brain neuroscientific research, there is a relative paucity of theoreti- activity during the meditative state as well as changes in neural cal reviews that consolidate the existing literature into a com- structure and function associated with meditation practice. To prehensive theoretical framework. our knowledge, no one has previously tried to consolidate the Existing research on mindfulness includes a few theoretical existing empirical literature. Our goal is to consolidate existing accounts describing mechanisms of mindfulness meditation findings and address several components that have been empiri- (see Table 1). Several of these accounts expound on the central cally supported. Furthermore, whereas the previous models role of attention in meditation practice (Brown & Ryan, 2003; have described the process of mindfulness almost exclusively Carmody, 2009; Lutz, Slagter, Dunne, & Davidson, 2008). from a conceptual, psychological perspective, this present Others have suggested that several components mediate the review will also integrate a neuroscientific perspective. This beneficial effects of mindfulness practice. For instance, Shap- review is meant not as a complete description of the research in iro, Carlson, Astin, and Freedman (2006) posit that attention, the field but to stimulate scientific debate. intention, and attitude are the three critical components of In this review, we first describe what we believe to be the mindfulness. Intentionally paying attention with a nonjudg- components of mindfulness meditation. We then discuss how mental attitude leads to a significant change in perspective, a these components are integrated during mindfulness medita- so-called decentering (Fresco et al., 2007) or reperceiving. tion and suggest how they might interact with each other dur- Brown, Ryan, and Creswell (2007) also describe several pro- ing a given situation. Finally, the relationship of self-compassion cesses underlying the beneficial effects of mindfulness, with the components is addressed.

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Table 2. Components Proposed to Describe the Mechanisms Through Which Mindfulness Works

Self-reported and experimental Mechanism Exemplary instructions behavioral findings Associated brain areas 1. Attention regulation Sustaining attention on the Enhanced performance: executive Anterior cingulate cortex chosen object; whenever attention (Attention Network distracted, returning attention Test and Stroop interference), to the object orienting, alerting, diminished attentional blink effect 2. Body awareness Focus is usually an object of Increased scores on the Observe Insula, temporo-parietal internal experience: sensory subscale of the Five Facet Mind- junction experiences of breathing, fulness Questionnaire; narrative emotions, or other body self-reports of enhanced body sensations awareness 3.1 Emotion regulation: Approaching ongoing emotional Increases in positive reappraisal (Dorsal) prefrontal cortex reappraisal reactions in a different way (Cognitive Emotion Regulation (PFC) (nonjudgmentally, with ac- Questionnaire) ceptance) 3.2 Emotion regulation: Exposing oneself to whatever is Increases in nonreactivity to inner Ventro-medial PFC, exposure, extinction, and present in the field of aware- experiences (Five Facet Mind- hippocampus, amygdala reconsolidation ness; letting oneself be affected fulness Questionnaire) by it; refraining from internal reactivity 4. Change in perspective on Detachment from identification Self-reported changes in self-con- Medial PFC, posterior the self with a static sense of self cept (Tennessee Self-Concept cingulate cortex, insula, Scale, Temperament and Char- temporo-parietal acter Inventory) junction

Components of Mindfulness Meditation the Theravada Buddhist tradition [Hart, 1987]; samatha in Tibetan Buddhist traditions [Lutz, Dunne, & Davidson, 2007]; We believe that an array of distinct but interacting mechanisms or dharana in Indian Yoga traditions [Vishnu Devananda, are at play in producing the benefits of mindfulness meditation 1999]). They often recommend a focused attention meditation practice and propose that the combination of the following before moving on to other types of meditations later in the components—some of which have been identified in previous learning process, such as those that focus on cultivating posi- accounts—describe much of the mechanism of action through tive emotions. In focused attention meditation ( HYPERLINK which mindfulness works: "" \l "bib148" \o "bib148" Lutz, Slagter, et al., 2008), attention is supposed to rest on a single object. Whenever the practitio- 1. Attention regulation ner notices that the mind has wandered off, she or he returns it 2. Body awareness to the chosen object. A typical instruction for a focused atten- 3. Emotion regulation, including tion meditation in the mindfulness meditation tradition is the a. Reappraisal following: “Focus your entire attention on your incoming and b. Exposure, extinction, and reconsolidation outgoing breath. Try to sustain your attention there without 4. Change in perspective on the self distraction. If you get distracted, calmly return your attention to the breath and start again” (Smith & Novak, 2003; p.77). These components interact closely to constitute a process Illustrating the effects of repeated practice of focused attention of enhanced self-regulation (Carver & Scheier, 2011; Vohs & meditation, meditators report that the regular practice enables Baumeister, 2004). However, the different components might them to focus their attention for an extended period of time come into play to varying degrees within any specific moment (Barinaga, 2003), and distractions disturb this focus less fre- during mindfulness meditation. In the following sections, we quently during formal meditation practice and in everyday address each of these components individually (see Table 2 for life. In accordance with such self-reports, a number of studies a list of the components and their characteristics). have empirically documented enhanced attentional perfor- mance in meditators (e.g., Jha et al., 2007; Slagter et al., 2007; Valentine & Sweet, 1999; van den Hurk, Giommi, Gielen, 1. Attention regulation Speckens, & Barendregt, 2010). Many meditation traditions emphasize the necessity to culti- Behavioral findings on meditation and executive attention. Dur- vate attention regulation early in the practice (e.g., samadhi in ing focused attention meditation, distracting external events as

Downloaded from pps.sagepub.com at The University of Iowa Libraries on May 24, 2012 540 Hölzel et al. well as memories or thoughts about future events represent con- identified when individuals engaged in a mindfulness prac- flicts to task goals. These are disregarded while the practitioner tice while awaiting unpleasant electric stimulation (Gard concentrates on the meditative object (e.g., the breath, body sen- et al., 2010). Five days of Integrative Body–Mind Training sations, thoughts, emotions, a mantra, or visualization). Main- may lead to greater activation of the rostral ACC during the taining the focus of attention on a pursued object, while resting state (Tang et al., 2009). Although ACC activation disregarding distractions, is referred to as conflict monitoring, might initially be enhanced when acquiring greater atten- or executive attention, and is one of the three attention networks tional control, it might later decrease with higher levels of proposed by Posner and Petersen (1990). One cognitive test that expertise, when the focus of attention is so steady that moni- specifically measures executive attention is the executive atten- toring distractions becomes superfluous (Brefczynski-Lewis, tion task of the Attention Network Test (Fan, McCandliss, Som- Lutz, Schaefer, Levinson, & Davidson, 2007). In addition to mer, Raz, & Posner, 2002). Two studies found that experienced these functional findings, structural MRI data also indicate meditators showed better performance on this executive atten- that meditation practice might exert an influence on the ACC. tion task when compared with nonmeditators, as indicated by Cortical thickness in the dorsal ACC was greater in experi- smaller error scores (Jha et al., 2007; van den Hurk et al., 2010) enced meditators compared with control subjects in an analy- and lower reaction times (Jha et al., 2007). Additionally, a lon- sis of brain gray matter (Grant, Courtemanche, Duerden, gitudinal study showed that only five days of meditation prac- Duncan, & Rainville, 2010), and 11 hr of Integrative Body– tice (Integrative Body–Mind Training) led to improvements on Mind Training led to an increase in white matter integrity in this test (Tang et al., 2007). Findings of the influence of mind- the ACC (Tang et al., 2010). In line with the assumption that fulness practice on executive attention using the classical Stroop ACC function is strengthened through concentrative medita- task (Stroop, 1935) are mixed. Whereas one study did not find tion, electroencephalogram data document increased frontal effects of an 8-week Mindfulness-Based Stress Reduction midline theta rhythm during meditation (Aftanas & Golo- course on Stroop interference (Anderson, Lau, Segal, & Bishop, cheikine, 2002; Kubota et al., 2001). Frontal midline theta is 2007), others using this test found lower Stroop interference in associated with attention demanding tasks and presumably experienced meditators compared with controls (Chan & Wool- reflects ACC (and medial prefrontal cortex) activity (Asada, lacott, 2007; Moore & Malinowski, 2009) and a reduction in Fukuda, Tsunoda, Yamaguchi, & Tonoike, 1999). Stroop interference following a brief meditation intervention Clinical relevance. The strengthening of attention regulation (Wenk-Sormaz, 2005). and accompanying ACC performance through mindfulness Neural mechanism of executive attention. Neuroimaging practice is especially promising for the treatment of disorders research has established that the anterior cingulate cortex that suffer from deficiencies in these functions, such as (ACC) enables executive attention (van Veen & Carter, 2002) attention-deficit/hyperactivity disorder (ADHD; e.g., Passa- by detecting the presence of conflicts emerging from incom- rotti, Sweeney, & Pavuluri, 2010) or bipolar disorder (Foun- patible streams of information processing. During meditation, toulakis, Giannakopoulos, Kovari, & Bouras, 2008). Although when distracting external events or memories conflict with there is currently insufficient evidence to support the effec- task goals, ACC activation may contribute to the maintenance tiveness of any type of meditation for ADHD (Krisanapra- of attention by alerting the systems implementing top-down kornkit, Ngamjarus, Witoonchart, & Piyavhatkul, 2010), regulation to resolve this conflict (van Veen & Carter, 2002). initial feasibility studies have shown promising effects on Together with the fronto-insular cortex, the ACC constitutes a improvements in attention (Zylowska et al., 2008). Bipolar network that is involved in switching between activations of disorder is also associated with impairments in sustained different brain networks, thereby facilitating cognitive control attention and executive function, as has been established by a (Sridharan, Levitin, & Menon, 2008). Neurons in these brain large number of empirical investigations (Ancin et al., 2010; regions have specific properties that enable a rapid relay of Clark, Iversen, & Goodwin, 2002; Kolur, Reddy, John, Kan- control signals to multiple areas of the brain (Allman, Watson, davel, & Jain, 2006; Kravariti et al., 2009; Maalouf et al., Tetreault, & Hakeem, 2005) to initiate responses during cogni- 2010). Neuroanatomical models of bipolar disorder propose a tively demanding tasks (Sridharan et al., 2008). key role of the ACC, and a meta-analysis has confirmed vol- Neuroscientific findings on meditation practice. Several neu- ume changes in the ACC as well as state-dependent alterations roscientific studies have reported the ACC to be implicated in resting state activity in this region (Fountoulakis et al., in meditation (Cahn & Polich, 2006). Using functional MRI 2008). Furthermore, ACC activation decreases during cogni- (fMRI), Hölzel et al. (2007) pursued the question of which tive tasks in bipolar patients (Gruber, Rogowska, & Yurgelun- brain region would be distinctly activated when meditators Todd, 2004). Mindfulness meditation practice might therefore performed focused attention meditation. Compared with be beneficial to ameliorate these deficits in cognitive function- age-, gender-, and education-matched controls, experienced ing and accompanying ACC function by strengthening these meditators showed greater activation in the rostral ACC skills in bipolar patients (Stange et al., in press). In line with (Hölzel et al., 2007), suggesting an effect of meditation this hypothesis, a few pilot studies have shown beneficial practice on ACC activity. A similar effect (greater rostral effects on symptoms in patients with bipolar disorder (Deck- ACC activation in meditators compared with controls) was ersbach et al., in press; Miklowitz et al., 2009; Williams et al.,

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2008). However, further research is needed to assess the effec- to Theravadan interpretations of Buddhist teachings, aware- tiveness of mindfulness-based treatments on attention regula- ness of the body was taught as the first frame of reference tion in these disorders. (foundation of mindfulness). In Thanissaro Bhikkhu’s transla- Effects of meditation practice on further components of atten- tion of the Satipatthana Sutta (Bhikkhu, 2010), the Buddha tion. Aside from the documented improvements in executive guides the monks with the following meditation instructions: attention through mindfulness, effects have also been reported In this way he [the monk] remains focused internally on the on other attention capacities. Within the framework of the net- body in and of itself, or externally on the body in and of itself, work components described by Posner and Petersen (1990), or both internally and externally on the body in and of itself. enhanced performance in “orienting” (directing and limiting Or he remains focused on the phenomenon of origination with attention to a subset of possible inputs) has been found follow- regard to the body, on the phenomenon of passing away with ing an 8-week mindfulness-based stress reduction course (Jha et regard to the body, or on the phenomenon of origination and al., 2007) and in experienced meditators, as compared with con- passing away with regard to the body. Or his mindfulness that trols (van den Hurk et al., 2010). An improvement in “alerting” “There is a body” is maintained to the extent of knowledge (achieving and maintaining a vigilant state of preparedness) was and remembrance. And he remains independent, unsustained found in experienced meditators following a 1-month mindful- by (not clinging to) anything in the world. This is how a monk ness retreat (Jha et al., 2007), as well as a 3-month samatha remains focused on the body in and of itself. retreat (MacLean et al., 2010). At a neurobiological level, these Self-report findings. Practitioners often self-report that the findings may relate to functional changes in the dorsal and ven- practice of attending to body sensations results in an enhanced tral attention systems (Corbetta & Shulman, 2002; Fox, Cor- awareness of bodily states and greater perceptual clarity of betta, Snyder, Vincent, & Raichle, 2006). These data suggest subtle interoception. In qualitative interviews, 10 experienced that early stages of mindfulness practice (represented by a short mindfulness meditators were asked what changes they experi- mindfulness course) may lead to improvement in the function of enced in their lives since they had begun meditating. Seven of the dorsal attention system involved in orienting, and more the 10 meditators spontaneously reported that they noticed a intensive open monitoring meditation on a 1-month retreat may more differentiated experience of body sensations, and four of additionally result in improvements in the function of the ven- them reported greater emotional awareness (Hölzel, Ott, Hem- tral attention system involved in alerting. pel, & Stark, 2006). Participants in a mindfulness-based stress Other types of attention tests have shown that 3 months of reduction course further illustrate self-reported changes in intensive mindfulness meditation lead to a smaller attentional body awareness, as measured by the Five Facet Mindfulness blink effect (a lapse in attention following a stimulus within a Questionnaire. Body awareness is represented in items of the rapid stream of presented stimuli) and modified distribution of Observe subscale of the questionnaire, which covers the brain resources (Slagter et al., 2007; also see van Leeuwen, awareness of body sensations (e.g., Item 1: “When I’m walk- Willer, & Melloni, 2009). Varying meditation practices will ing, I deliberately notice the sensations of my body moving”), differentially affect these specific attentional components (see hearing, smelling, seeing, interoceptions, thoughts, and emo- Lutz, Slagter, et al., 2008, for a review of focused attention tions (e.g., Item 11: “I notice how foods and drinks affect my versus open monitoring meditation). Future research is neces- thoughts, bodily sensations, and emotions”). Participation in sary to assess the impact of different types of meditation prac- the mindfulness-based stress reduction course resulted in large tice on these other attentional components. increases in scores on this scale (Carmody & Baer, 2008). Attention regulation—in particular conflict monitoring— Behavioral findings. Although meditation practitioners report seems to be an important mechanism that is often developed improved capability for body awareness, to our knowledge early in mindfulness practice. A sufficient degree of attention there has been no empirical evidence to verify these claims. In regulation is necessary in order to stay engaged in meditation, fact, studies that tested this claim by assessing performance as opposed to drifting off into day dreaming. Thus, successful on a heartbeat detection task, a standard measure of resting attention regulation might be a building block for practitioners interoceptive awareness, found no evidence that meditators to also benefit from the other mechanisms of mindfulness had superior performance compared with nonmeditators practice, which will be described below. This connection (Khalsa et al., 2008; Nielsen & Kaszniak, 2006). However, between attention regulation and other mechanisms should be awareness of heartbeat sensations is not emphasized during tested in future research. mindfulness practice and thus may not be the best index of the awareness cultivated by the practice of meditation. Further studies are needed that test other types of body awareness, 2. Body awareness such as tactile acuity, which has been shown to be superior in Body awareness can be understood as the ability to notice experienced Tai Chi practitioners compared with matched subtle bodily sensations (Mehling et al., 2009). In mindfulness controls (Kerr et al., 2008). practice, the focus of attention is usually an object of internal Neuroscientific findings: Functional neuroimaging. A number of experience: sensory experiences of breathing, sensory experi- findings from the mindfulness neuroscience literature point to ences related to emotions, or other body sensations. According changes in the function and structure of brain regions related

Downloaded from pps.sagepub.com at The University of Iowa Libraries on May 24, 2012 542 Hölzel et al. to body awareness. The insula is commonly activated during own emotional life; in turn, an awareness of one’s emotions is tasks of interoceptive awareness (Craig, 2003), and its local a precondition for being able to regulate those emotions. Help- gray matter volume correlates with interoceptive accuracy and ing individuals increase their body awareness can therefore be visceral awareness (Critchley, Wiens, Rotshtein, Ohman, & considered a relevant aspect in the treatment of psychological Dolan, 2004). Insula activation has been found to be increased disorders. For example, a lack of awareness of internal experi- in individuals after a mindfulness-based stress reduction ence—along with problems in emotion regulation—is a cru- course (compared with individuals who had not practiced cial problem for individuals with borderline personality mindfulness) when they focused on their momentary experi- disorder, and helping patients increase their internal awareness ence (i.e., employed an experiential focus; Farb et al., 2007). might be one key element in its treatment (Linehan, Arm- This study also found increased activation of the secondary strong, Suarez, Allmon, & Heard, 1991; Wupperman, Neu- somatosensory area, which is relevant for the processing of mann, & Axelrod, 2008). Furthermore, the increase of body exteroceptive sensory events. In another study, a group of par- awareness is also relevant in the treatment of eating disorders ticipants that had undergone mindfulness training showed (Hill, Craighead, & Safer, 2011) as well as substance abuse greater activation of the right insula when being presented disorders. In a pilot study of 16 heroin users in early recovery, with sad movie clips (Farb et al., 2010). Further neuroscien- high levels of the Observe subscale of the Five Facet Mindful- tific evidence along the same lines comes from studies on ness Questionnaire were associated with decreased heroin use mindfulness in the context of pain. When presented with among those at high risk for relapse (Schuman-Olivier, Alba- unpleasant stimuli during a mindful state, mindfulness medita- nese, Carlini, & Shaffer, 2011), suggesting a role for body tors show stronger brain activation in the (posterior) insula and awareness in the recovery process. secondary somatosensory cortex (Gard et al., 2010). Similarly, Body awareness and empathy. Internal awareness of one’s mindfulness meditators more robustly activated the left ante- own experience has also been suggested to be an important pre- rior, posterior, and mid-insula as well as the thalamus (Grant, condition for empathic responses. Accurate observations of the Courtemanche, & Rainville, 2010). The enhanced sensory self are required for the appropriate understanding of others processing has been suggested to represent increased bottom- (Decety & Jackson, 2004). Self-report studies provide empirical up processing of the stimulus, that is, awareness of the actual support for the existence of this relationship. A higher level of sensation of the stimulus as it is. mindful observation, as assessed with the Observe scale of the Neuroscientific findings: Structural neuroimaging. Two cross- Kentucky Inventory of Mindfulness Skills (Baer, Smith, & sectional studies comparing the gray matter morphometry of Allen, 2004), has been found to be associated with more engage- the brains of experienced meditators and controls showed that ment in empathy (Dekeyser, Raes, Leijssen, Leysen, & Dewulf, meditators had greater cortical thickness (Lazar et al., 2005) 2008), as assessed with the Interpersonal Reactivity Index and greater gray matter concentration (Hölzel et al., 2008) in (Davis, 1980). Neuroscientific research shows that a subset of the right anterior insula. Although 8 weeks of mindfulness brain regions (namely, the insula and temporo-parietal junction) practice did not reveal changes in gray matter concentration in is impacted both in awareness of one’s own body sensations and the insula (Hölzel et al., 2011), the same study did reveal that in social cognition and empathic responses (Singer et al., 2004). 8 weeks of practice led to increases in gray matter concentra- Enhanced function of these structures following mindfulness tion in the temporo-parietal junction. It has been suggested training might therefore also correspond to improved empathic that the temporo-parietal junction is a crucial structure for responses and compassion attributed to meditation training mediating the first-person perspective of bodily states (Blanke (Shapiro, Schwartz, & Bonner, 1998). Supporting this assump- et al., 2005), or embodiment (Arzy, Thut, Mohr, Michel, & tion, research has found that Tibetan monks with over 10,000 hr Blanke, 2006), and that impaired processing at the temporo- of meditation experience showed greater activation of both parietal junction may lead to the pathological experience of regions during compassion meditation (a meditation that aims at the self, such as out-of-body experiences (Blanke & Arzy, cultivating feelings of empathy toward the suffering of other 2005). Morphological changes in the temporo-parietal junc- beings and the wish to alleviate their suffering) while they were tion might be associated with an increased awareness of the presented with auditory stimuli of people suffering (Lutz, Bref- experience of oneself within the body. Such changes seem to czynski-Lewis, Johnstone, & Davidson, 2008). correspond with translations of meditation instructions To summarize, body sensations are a common object of ascribed to the historical Buddha (Bhikkhu, 2010): “His mind- attention during mindfulness meditation, and practitioners fulness that ‘There is a body’ is maintained to the extent of report improved body awareness. Although there have been no knowledge and remembrance.” objective behavioral data supporting the increased awareness, Body awareness and emotion regulation. Body sensations neuroscientific data on mindfulness practice point to the modi- have been ascribed a crucial role in the conscious experience fication of brain regions involved in first-person conscious of emotions (feelings), not only historically (James, 1884), but experience of body awareness. The enhancement of body also currently (Bechara & Naqvi, 2004; Damasio, 1999, 2003). awareness might have relevance for affect regulation and An increased awareness of the body’s response to an emo- empathic processes and thus may be particularly relevant in tional stimulus might thus lead to greater awareness of one’s the mindfulness-based treatment of patients with such deficits.

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Ongoing attempts to advance the development of instruments Physiological studies also support the proposition that for the valid assessment of body awareness (Mehling et al., meditation training leads to decreased emotional reactivity 2009) will help to further illuminate this connection. and facilitates a return to emotional baseline after reactivity. For example, experienced mindfulness meditators have shown a faster decrease in skin conductance in response to 3. Emotion regulation aversive stimuli (Goleman & Schwartz, 1976), as well as less In this section, we first describe the general findings regard- enhancement of the startle response by aversive stimuli ing the effects of mindfulness on emotion regulation. Then (Zeidler, 2007). An electroencephalogram study found that we focus on two different emotion regulation strategies that mindfulness-based stress reduction training led to increases in seem to be involved in mindfulness, namely, reappraisal and left-sided anterior brain activation after the course compared extinction. with a wait-list control group (Davidson et al., 2003). This pat- Role of emotion regulation in meditation practice. A growing tern of lateralization has previously been associated with the body of literature suggests that mindfulness practice results in experience of positive emotions (Davidson, 1992). Similarly, improvements in emotion regulation. Emotion regulation stronger relative left prefrontal activation was also recently refers to the alteration of ongoing emotional responses through found as a state effect in previously depressed individuals fol- the action of regulatory processes (Ochsner & Gross, 2005). In lowing a short practice of mindful breathing and loving kind- Theravadan translations of Buddhist teachings, the alteration ness meditation (Barnhofer, Chittka, Nightingale, Visser, & of emotional responses is addressed in the Satipatthana Sutta Crane, 2010). These findings support the proposition that in terms of the overcoming of sorrow and distress as part of mindfulness practice has an effect on the physiological aspects meditation practice: of positive emotions and thus positively influences emotional processing. This is the direct path for the purification of beings, for the overcoming of sorrow and lamentation, for the disap- Neural mechanisms of emotion regulation. During emo- pearance of pain and distress, for the attainment of the tion regulation, prefrontal control systems modulate emotion- right method, and for the realization of Unbinding. . . . He generative systems, such as the amygdala, which is responsible [the monk] remains focused on feelings . . . mind . . . for the detection of affectively arousing stimuli (Ochsner & mental qualities in and of themselves—ardent, alert, and Gross, 2005). More specifically, these prefrontal structures mindful—putting aside greed and distress with reference include dorsal regions of the lateral prefrontal cortex (PFC) to the world. (Bhikkhu, 2010) that have been implicated in selective attention and working memory; ventral parts of the PFC implicated in response inhi- Behavioral and peripheral physiological finding. Studies bition; the ACC, which is involved in monitoring control pro- from the field of mindfulness research have addressed cesses; and the dorso-medial PFC implicated in monitoring improvements in emotion regulation through a variety of one’s affective state (Modinos, Ormel, & Aleman, 2010; Och- approaches, including experimental, self-report, peripheral sner & Gross, 2008). A typical pattern detected when individu- physiological, and neuroimaging data. Healthy novices als deliberately regulate affective responses is increased enrolled in a 7-week mindfulness training program showed a activation within the PFC and decreased activation in the reduction in emotional interference (assessed as the delay in amygdala (Beauregard, Levesque, & Bourgouin, 2001; Haren- reaction time after being presented with affective versus neu- ski & Hamann, 2006; Schaefer et al., 2002), suggesting that tral pictures) compared with those who followed a relaxation PFC projections to the amygdala exert an inhibitory top-down meditation protocol and those in a wait-list control group influence (Banks, Eddy, Angstadt, Nathan, & Phan, 2007; (Ortner et al., 2007). Both the mindfulness and relaxation Davidson, Jackson, & Kalin, 2000).2 meditation groups also displayed significant reductions in physiological reactivity during the task. Furthermore, in a Psychological disorders and emotion regulation. A variety group of long-term practitioners, participants with more mind- of psychological disorders are associated with reduced emo- fulness meditation experience showed less emotional interfer- tion regulation capacity (Cicchetti, Ackerman, & Izard, 1995; ence than did less experienced practitioners. Studies using Davidson, 2000; Gross, 1998; Hayes, Wilson, Gifford, Fol- self-report data from healthy individuals have shown that lette, & Strosahl, 1996; Mennin, Heimberg, Turk, & Fresco, mindfulness meditation decreased negative mood states (Jha, 2002). Disorders characterized by a deficit in emotion regula- Stanley, Kiyonaga, Wong, & Gelfand, 2010), improved posi- tion are frequently associated with dysfunction in the frontal- tive mood states, and reduced distractive and ruminative limbic network, that is, reduced prefrontal activation and thoughts and behaviors (Jain et al., 2007). A questionnaire exaggerated amygdala activation (e.g., depression, H. C. study investigating the immediate effects of brief (15-min) Abercrombie et al., 1998; borderline personality disorder, Sil- stress management interventions found that mindful breathing bersweig et al., 2007; bipolar disorder, Pavuluri, O’Connor, may help to reduce reactivity to repetitive thoughts (Feldman, Harral, & Sweeney, 2007; social phobia, Phan, Fitzgerald, Greeson, & Senville, 2010). Nathan, & Tancer, 2006; obsessive-compulsive disorder,

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Breiter & Rauch, 1996; posttraumatic stress disorder, Shin et al., regulation strategies that might be influenced by mindfulness 2005; impulsive aggression, Coccaro, McCloskey, Fitzgerald, & practice. Phan, 2007; addiction, Bechara, 2005; generalized anxiety, Monk There are several proposed classifications for different et al., 2008; and trait anxiety, Etkin et al., 2004). kinds of emotion regulation (Ochsner & Gross, 2005; Parkin- son & Totterdell, 1999). Ochsner and Gross (2005) have sug- Neuroscientific findings on mindfulness and emotion reg- gested a distinction between behavioral regulation (e.g., ulation. In contrast to psychiatric disorders that are character- suppressing expressive behavior) and cognitive regulation. ized by suboptimal or deficient emotion regulatory responses Cognitive regulation can rely on attentional control (e.g., and corresponding abnormalities in brain activation patterns, a selective inattention to emotional stimuli, performing distract- few neuroimaging studies have found increased prefrontal ing secondary tasks) or on cognitive change. Cognitive change activation and improved prefrontal control over amygdala strategies include the controlled regulation of an ongoing responses in association with mindfulness. The earliest evi- emotional response, such as reappraisal (i.e., reinterpreting dence for the neurophysiological basis of differences in emo- the meaning of a stimulus to change one’s emotional response tion regulation according to dispositional levels of mindfulness to it) and extinction (stimulus-response reversal). As discussed was found in a study that investigated the effect of disposi- in Section 1, attentional control plays a crucial role in mindful- tional (trait) mindfulness on brain activation while healthy ness meditation. Whereas typical contemporary descriptions participants labeled the affect of emotional facial expressions regard attentional control in emotion regulation as adaptive (Creswell, Way, Eisenberger, & Lieberman, 2007). Higher dis- when attention is directed away from emotionally distressing positional mindfulness, as measured by the Mindful Attention material, mindfulness usually involves bringing attention to Awareness Scale, predicted (a) increased activation at multiple the stimulus. Keeping attention on an emotional reaction leads sites of the prefrontal cortex (including the ventromedial PFC, to a situation of exposure with a subsequent extinction pro- medial PFC, and ventrolateral PFC), (b) reduced amygdala cess. Extinction plays a crucial role in producing the beneficial activity, and (c) a stronger inhibitory association between effects of mindfulness meditation and will be discussed in amygdala activity and regions of the PFC. detail below. The following section explores the role of Although the above mentioned study investigated disposi- reappraisal. tional mindfulness in individuals without any mindfulness training, other studies have reported evidence that mindful- Reappraisal. Reappraisal has been suggested to be one of the ness meditation involves activation of brain regions relevant ways in which emotion gets regulated during mindfulness. for emotion regulation, and thus activation of these regions Garland et al. (2011) described mindful emotion regulation as might be modified through mindfulness practice. During “positive reappraisal,” or the adaptive process through which mindfulness meditation, experienced mindfulness meditators stressful events are reconstrued as beneficial, meaningful, or show greater activation in the dorso-medial PFC and rostral benign (e.g., thinking that one will learn something from a dif- ACC compared with nonmeditators (Hölzel et al., 2007). After ficult situation). A very recent self-report study showed that participants completed an 8-week mindfulness-based stress mindfulness practice leads to increases in positive reappraisal reduction course, Farb et al. (2007) found increased activity in and that these increases mediate an improvement in stress lev- participants’ ventrolateral PFC, which the authors interpreted els (Garland et al., 2011). as augmented inhibitory control. Following participation in a mindfulness-based stress reduction course, social anxiety Neuroscientific findings. Cognitive reappraisal of aversive patients presented with negative self-beliefs showed a quicker stimuli has been found to coincide with activity in the dorso- decrease of activation in the amygdala as compared with mea- lateral PFC, orbitofrontal PFC, and ACC (Eippert et al., 2007; sures taken before course completion (Goldin & Gross, 2010). Ochsner, Bunge, Gross, & Gabrieli, 2002; Ochsner et al., The observed improvements in emotion regulation associ- 2004), and dorsal PFC activity has been found to go along ated with mindfulness practice likely underlie many of the posi- with reappraisal success (Modinos et al., 2010; Wager, David- tive effects of mindfulness practice on mental health. Indeed, son, Hughes, Lindquist, & Ochsner, 2008). Thus, whereas improved emotion regulation underlies the beneficial effects of extinction processes may depend more upon ventral frontal mindfulness practice on stress reduction (Garland, Gaylord, & systems (such as the ventromedial PFC) that are directly con- Fredrickson, 2011) and on reductions of depressive symptoms nected with the subcortical systems (see Exposure, extinction, (Shahar, Britton, Sbarra, Figueredo, & Bootzin, 2010). and reconsolidation for details), reappraisal may depend more on dorsal frontal systems (Ochsner & Gross, 2008). Different strategies of emotion regulation. Although it To investigate the neural correlates of dispositional mind- seems well established that mindfulness has positive effects on fulness in the context of reappraisal, Modinos et al. (2010) emotion regulation, the exact processes underlying these assessed fMRI study participants’ individual differences in improvements seem less clear. Emotion regulation is an dispositional mindfulness with the Kentucky Inventory of umbrella term for a wide array of strategies for altering emo- Mindfulness Skills and asked the participants to either attend tional responses. Here, we will consider some emotion to or reappraise negative pictures. Findings showed that levels

Downloaded from pps.sagepub.com at The University of Iowa Libraries on May 24, 2012 How Does Mindfulness Meditation Work? 545 of dispositional mindfulness were positively correlated with “positive reappraisal,” and correspondingly, studies find activations in the left and right dorsomedial PFC during the enhanced brain activity in multiple prefrontal regions involved reappraisal condition. Therefore, trait mindfulness seems to be in cognitive change strategies. However, other studies have positively associated not only with reappraisal success but conceptualized the changes as “nonappraisal” and have identi- also with increased activation in brain regions that support this fied decreased brain activity in prefrontal regions. Further kind of emotion regulation. research is needed to test the hypothesis that the amount of required prefrontal control decreases with increased expertise. Does mindfulness involve reappraisal or nonappraisal? In the following section, we will turn toward exposure, extinc- Although the above findings suggest increased “reappraisal” tion, and reconsolidation as a further mechanism of action of related to mindfulness (Garland et al., 2011), there also seems mindfulness meditation. to be some inconsistency in the literature. Other work has identified decreased cognitive control associated with mind- Exposure, extinction, and reconsolidation. During mind- fulness meditation, interpreted as nonappraisal. As mentioned fulness, practitioners expose themselves to whatever is present earlier in the section on body awareness, experienced medita- in the field of awareness, including external stimuli as well as tors, when presented with unpleasant or painful stimuli during body sensations and emotional experiences. They let them- a mindful state (Gard et al., 2010) or a baseline state (Grant, selves be affected by the experience, refraining from engaging Courtemanche, & Rainville, 2011) showed enhanced sensory in internal reactivity toward it, and instead bringing accep- processing, that is, increased bottom-up processing of the tance to bodily and affective responses (Hart, 1987). Practitio- stimulus. At the same time, decreased prefrontal activation ners are instructed to meet unpleasant emotions (such as fear, was observed in meditators in both studies when they were sadness, anger, and aversion) by turning towards them, rather presented with the painful stimuli. These findings were than turning away (Santorelli, 2000). Those people who are explained with a decrease in top-down control, representing a new to meditation often initially find this process counterintui- lack of reappraisal. tive, but many practitioners discover that the unpleasant emo- This discrepancy brings to light a question: Does emotion tions pass away and a sense of safety or well-being can be regulation during mindfulness involve cognitive control (and experienced in their place. corresponding prefrontal engagement), or is it characterized Parallels between the process described here and exposure instead by its absence? Whereas the acceptance of one’s emo- therapy are evident. Exposure therapy is a highly effective tional response is characterized by the absence of active cogni- behavioral therapy technique for reducing fear and anxiety tive control over the emotional reaction, bringing mindful responses (Chambless & Ollendick, 2001). Its core element is awareness to emotional responses might initially require some to expose patients to fear-provoking stimuli and prevent their cognitive control, in order to overcome habitual ways of inter- usual response in order for them to extinguish the fear response nally reacting to one’s emotions. Although currently specula- and to instead acquire a sense of safety in the presence of the tive, it seems possible that the degree of meditation expertise formerly feared stimuli (Öst, 1997). Clinical studies on expo- of the individual might be relevant when considering the ques- sure therapy show that access to safety behaviors can interfere tion of whether mindfulness involves cognitive control or its with the beneficial effects of an exposure situation (Lovibond, absence. Whereas beginners might require more active cogni- Mitchell, Minard, Brady, & Menzies, 2009; Salkovskis, Clark, tive regulation in order to approach ongoing emotional reac- Hackmann, Wells, & Gelder, 1999; Wells et al., 1995). Safety tions in a different way and might therefore show greater behaviors include not only overt behavior (such as avoiding prefrontal activation, expert meditators might not employ this eye contact in social phobia) but also cognitive avoidance. prefrontal control. Rather, they might use different strategies; Mindfulness meditation includes refraining from engaging in they may have automated an accepting stance toward their cognitive avoidance or other safety behaviors by using experience so they no longer require cognitive control efforts, enhanced attention regulation skills, thereby maximizing the and they could have different baseline blood flow as a conse- exposure to the experienced emotion. quence of plastic processes. A similar interpretation has previ- Additionally, meditation is often associated with high lev- ously been suggested by Brefczynski-Lewis et al. (2007) in els of relaxation in the form of increased parasympathetic tone regard to attentional control. and decreased sympathetic activity (Benson, 2000). Peripheral To summarize, several studies have demonstrated improve- physiological changes have been observed with some consis- ments in emotion regulation associated with mindfulness. Psy- tency (but see Shapiro, 1982), including decreased heart rate chological disorders characterized by problems in emotion (Zeidan, Johnson, Gordon, & Goolkasian, 2010), decreased regulation, such as mood disorders, anxiety disorders, or bor- blood pressure (de la Fuente, Franco, & Salvator, 2010), derline personality disorder, can benefit from the enhancement decreased cortisol levels (Carlson et al., 2007), decreased of emotion regulation capacities. Different emotion regulation breathing rate (Lazar et al., 2005), lowered oxygen and carbon strategies might show improvements following mindfulness dioxide consumption (Young & Taylor, 1998), decreased skin practice. Some studies have conceptualized the improved conductance response (Austin, 2006), and decreased muscle emotion regulation associated with mindfulness practice as tension (Benson, 2000). Since extinction mechanisms are

Downloaded from pps.sagepub.com at The University of Iowa Libraries on May 24, 2012 546 Hölzel et al. thought to be supported by the experience of a state of relax- 2005) positively correlated with extinction recall. In addition, ation while the individual encounters the feared stimuli hippocampal activation has also been found to be involved in (Wolpe, 1958), the relaxation component of meditation might fear extinction recall. Functional connectivity analysis reveals serve to maximize the effects of the extinction process. that the vmPFC and hippocampus work in concert during In the Five Facet Mindfulness Questionnaire, the capacity extinction recall to inhibit fear, suggesting that they comprise to expose oneself to internal experience without reactivity is a network that mediates the expression of extinction memory captured in the Non-Reactivity to Inner Experience Scale. in the appropriate context (Milad et al., 2007). Hippocampal Example items are “In difficult situations, I can pause without activation during extinction recall is likely related to signaling immediately reacting” (Item 21), or “When I have distressing the extinguished context (contextual safety; Corcoran, Des- thoughts or images I am able just to notice them without react- mond, Frey, & Maren, 2005; Corcoran & Maren, 2001). The ing” (Item 29). With completion of a mindfulness-based stress amygdala has been implicated in both human and animal stud- reduction course, scores on this scale increase with large effect ies as playing a crucial role during the acquisition and expres- sizes (Carmody & Baer, 2008), corroborating the theory that sion of conditioned fear (Davis & Whalen, 2001; LeDoux, mindfulness practice leads to the self-perception of decreased 2000; Pare, Quirk, & Ledoux, 2004; Phelps & LeDoux, 2005), reactivity. This likely is a mechanism for facilitating including the detection of stressful and threatening stimuli and exposure. the initiation of adaptive coping responses (Hasler et al., 2007). When individuals regulate their emotions, the amyg- Fear conditioning, extinction, and reconsolidation. The pro- dala is thought to be down-regulated by the vmPFC and hip- cess of fear extinction has been studied extensively in the con- pocampus (Banks et al., 2007; Davidson et al., 2000; Milad, text of conditioned fear. Fear conditioning is a learning process Rauch, Pitman, & Quirk, 2006), both of which have extensive in which a neutral conditioned stimulus (e.g., a tone) is paired connections with the amygdala. This inhibition of the amyg- with an aversive unconditioned stimulus (e.g., a shock). After a dala serves to suppress fear (Milad et al., 2006; Rauch, Shin, few pairings, the presentation of the conditioned stimulus comes & Phelps, 2006), thereby allowing control over behavioral to also elicit various fear responses (e.g., freezing in animals; reactions to emotions (Price, 2005). Deficits in fear extinction sympathetic arousal in humans). Repeated presentations of the are thought to be related to a number of psychiatric disorders, conditioned stimulus in the absence of the unconditioned stimu- and neuroimaging studies have shown that the aforementioned lus result in the extinction of the conditioned responses. Extinc- structures are dysfunctional in several psychiatric disorders, tion does not erase the initial association between conditioned such as posttraumatic stress disorder (Milad et al., 2009), and unconditioned stimuli but is thought to form a new memory schizophrenia (Holt et al., 2009), and depression (Anand et al., trace (Quirk, 2002; Rescorla, 2001) or reconsolidate the old 2005). memory with new contextual associations (Inda, Muravieva, & Alberini, 2011; Nader & Einarsson, 2010; Rossato, Bevilaqua, Effects of meditation practice on the neural network Izquierdo, Medina, & Cammarota, 2010). After extinction train- underlying extinction. There is recent evidence from ana- ing, extinction memory is thought to compete with conditioned tomical MRI studies that the aforementioned brain regions memory for control of fear expression (Myers & Davis, 2007). show structural changes following mindfulness meditation Recent research has shown that successful extinction memory training. Cross-sectional studies comparing mindfulness med- reliably differentiates healthy from pathological conditions itators and nonmeditators found that meditators showed (Holt et al., 2009; Milad et al., 2008). Extinction learning and its greater gray matter concentration in the hippocampus (Hölzel retention may thus be a critical process in the transformation of et al., 2008; Luders, Toga, Lepore, & Gaser, 2009). Further- maladaptive states. It allows individuals to learn not to have a more, Hölzel et al. (2011) recently observed that structural fear response to neutral stimuli, when there is no adaptive func- changes in the hippocampus were detectable within a period tion for the fear response. Rather, individuals can flexibly elicit of only 8 weeks in participants that underwent mindfulness- other more adaptive emotional and behavioral responses. based stress reduction, and Hölzel et al. (2008) found that cumulative hours of meditation training were positively Neural mechanisms of fear extinction and extinction correlated with gray matter concentration in the vmPFC in retention. Recent fMRI research on fear conditioning has experienced meditators. In a longitudinal study enrolling par- identified a network of brain regions that are crucial for the ticipants in an 8-week mindfulness-based stress reduction extinction of conditioned fear responses and its retention. As course, Hölzel et al. (2010) found an impact of the stress- outlined below, this network seems to strengthen through reducing effects of mindfulness meditation on the amygdala; mindfulness practice. The ventromedial prefrontal cortex the greater the decrease in participants’ scores on perceived (vmPFC) has been shown to be important for the successful stress over the 8 weeks, the greater a decrease they showed in recall of the extinction (Milad & Quirk, 2002; Morgan, gray matter concentration in the right amygdala. Modified Romanski, & LeDoux, 1993; Quirk, Russo, Barron, & Lebron, gray matter concentration in these regions that is dependent on 2000), with the magnitude of vmPFC activation (Milad et al., meditation training might potentially be related to the improved 2007) and the cortical thickness of the vmPFC (Milad et al., ability to regulate emotional responses. Furthermore, fMRI

Downloaded from pps.sagepub.com at The University of Iowa Libraries on May 24, 2012 How Does Mindfulness Meditation Work? 547 studies show that meditation involves activation of the hippo- (Olendzki, 2010). When internal awareness becomes enhanced campus and medial PFC (Lazar et al., 2000; Lou et al., 1999; through meditation, meditators report that they can observe Newberg et al., 2001), suggesting that regular meditation prac- mental processes with increasing clarity (cf. MacLean et al., tice enhances the function of these brain regions. Additionally, 2010) and increasing temporal resolution. Within this enhanced for those with social anxiety disorder, amygdala activation is clarity, the process of a repeatedly arising sense of self becomes reduced following 8 weeks of mindfulness practice (Goldin & observable to the meditator through development of meta- Gross, 2010). There thus appear to be striking similarities awareness. Meta-awareness is a form of subjective experience in the brain regions being influenced by mindfulness medita- and executive monitoring, in which one takes a nonconceptual tion and those involved in mediating fear extinction. These perspective as a distributed form of attention toward the con- findings suggest that mindfulness meditation could directly tents of conscious experience and the processes involved. influence one’s capacity to extinguish conditioned fear by Meta-awareness is not entangled in the contents of awareness enhancing the structural and functional integrity of the brain (Deikman, 1982; Raffone & Pantani, 2010; Varela, Thompson, network involved in safety signaling. The neuroscientific con- & Rosch, 1991) and facilitates a detachment from identifica- siderations described here support the previously held view tion with the static sense of self. Rather than as a static entity that extinction might contribute to some of the beneficial or structure, the sense of self can be experienced as an event effects of mindfulness practice (Baer, 2003; Brown et al., (Olendzki, 2006). It has been postulated that paying close 2007). attention to the transitory nature of this sense of self leads to the “deconstruction of the self” (Epstein, 1988). The Dalai The impact of extinction processes within meditation Lama describes the resulting understanding that practitioners practice. The role of extinction processes in the improve- reach: “This seemingly solid, concrete, independent, self- ments following mindfulness-based treatments is most obvi- instituting I under its own power that appears actually does not ous in the treatment of anxiety disorders, which have reliably exist at all” (Gyatso, 1984, p.70). In place of the identification been found to benefit from mindfulness practice (Kabat-Zinn with the static self, there emerges a tendency to identify with et al., 1992; Kimbrough, Magyari, Langenberg, Chesney, & the phenomenon of “experiencing” itself.3 Berman, 2010; Roemer et al., 2008). Nonreactivity and the From a Buddhist perspective, identification with the static successive extinction mechanism presumably also play a cru- sense of self is the cause of psychological distress, and dis- cial role in the stress-reducing effects of mindfulness and identification results in less afflictive experience and the free- might mediate decreased perceived stress scores (Carmody & dom to experience a more genuine way of being (Olendzki, Baer, 2008; Chang et al., 2004). They might also be highly 2010). As the psychologist Jack Engler (2004) puts it: relevant for the benefits of mindfulness in the treatment of When it is realized that no self is to be found in the ele- substance abuse (Brewer et al., 2009). Beyond that, exposure ments of our experience, it begins the process of liberation. is pursued toward whatever emotions present themselves, Understanding that our sense of “I” is not as solid, permanent, including sadness, anger, and aversion, as well as pleasant or substantial as we habitually hold it to be ultimately uproots emotions, such as happiness. We therefore suggest that extinc- clinging, attachment, and hostility. Understanding this burns tion is effective during all of these emotional experiences, up the fuel that runs our repetitive habits. Those who have leading to an overwriting of previously learned stimulus- understood this report a sense of spacious lightness and free- response associations. Buddhist teachings claim that the non- dom. They exhibit deep concern and tenderness for others. clinging to unpleasant and pleasant experiences leads to According to Buddhist philosophy, a change in perspective liberation (Olendzki, 2010). Framed in Western psychological on the self is thus the key in the process to enduring forms of terminology, one could say that nonreactivity leads to unlearn- happiness. ing of previous connections (extinction and reconsolidation) Whereas more advanced meditation practices are required and thereby to liberation from being bound to habitual emo- to experience this drastic disidentification from the static sense tional reactions. of self, a de-identification from some parts of mental content is often experienced even in the earliest stages of meditation practice. In mindfulness practice, all experiences are observed 4. Change in perspective on the self as they arise and pass. By closely observing the contents of The essence of Buddhist psychology lies in the teaching that consciousness, practitioners come to understand that these are there is no such thing as a permanent, unchanging self (Olen- in constant change and thus are transient. The mindful, non- dzki, 2010). Rather, the perception of a self is a product of an judgmental observation fosters a detachment from identifica- ongoing mental process. This perception reoccurs very rapidly tion with the contents of consciousness. This process has been in the stream of mental events, leading to the impression that termed “reperceiving” or “decentering” (Carmody, Baer, the self is a constant and unchanging entity. The self is experi- Lykins, & Olendzki, 2009; Fresco et al., 2007; Shapiro et al., enced as being the one who inhabits the body, being the one 2006) and has been described as the development of the who is thinking the thoughts, being the one experiencing emo- “observer perspective” (Kerr, Josyula, & Littenberg, 2011). tions, and being the agent of actions; having free will We suggest that although this stage is not yet the full

Downloaded from pps.sagepub.com at The University of Iowa Libraries on May 24, 2012 548 Hölzel et al. disidentification from a static and unchanging self described including areas of the medial PFC (Gusnard, Akbudak, Shul- above, it is a change in perspective about the sense of self and man, & Raichle, 2001; Kelley et al., 2002; Sajonz et al., 2010), an alteration in first-person subjective experience. posterior cingulate cortex /anterior precuneus (Sajonz et al., Philosophical considerations, theoretical accounts, and 2010), and also the inferior parietal lobule (Sajonz et al., experiential reports ascribe to the change in the perspective on 2010). The medial PFC has been shown to support an array of the self a crucial role for development and maturity in medita- self-related capacities, including memory for self-traits (Kel- tion. However, perspective on the self is rather difficult to ley et al., 2002; Macrae, Moran, Heatherton, Banfield, & Kel- operationalize, and little empirical research has been pub- ley, 2004) or reflected self-knowledge (Lieberman, Jarcho, & lished that documents these types of changes following mind- Satpute, 2004). The posterior cingulate cortex and precuneus fulness meditation. This area of research could benefit from are engaged when individuals assess the relevance or signifi- the introduction of clear and consistent definitions of self- cance of a stimulus for themselves (Gusnard et al., 2001; related processes as well as terms that until now have been Schmitz & Johnson, 2007) and have been suggested to be par- applied inconsistently among various authors and disciplines ticularly important for the integration of self-referential stim- (e.g., I, me, ego, self, etc.). Although a clarification of the defi- uli in the emotional and autobiographical context of one’s own nitions, theories, and conceptualization is far beyond the scope person (Northoff & Bermpohl, 2004). All of these structures of this article and is not its focus (but see Legrand & Ruby, show high activity during rest, mind wandering, and condi- 2009, and Strawson, 2000), we summarize the few self-report tions of stimulus-independent thought (Northoff et al., 2006) and neuroimaging studies that touch on a change in the per- and have therefore also been characterized as the “default spective on the self through mindfulness practice. mode” of the brain or as the default mode network (Buckner, Self-report findings. Self-report studies have begun to docu- Andrews-Hanna, & Schacter, 2008; Gusnard & Raichle, ment the experienced changes in perspective on the self fol- 2001). lowing mindfulness practice. In a qualitative analysis of A few MRI studies have begun looking at activity in the diaries, Kerr and colleagues focused on the development of an default mode network in association with mindfulness prac- “observing self,” or meta-perspective on experience, and have tice. Comparing brain activation during mindfulness medita- described how participants experience this shift over the tion versus a resting state reveals decreased brain activity in course of an 8-week mindfulness-based stress reduction course subsystems of the default mode network (Ott, Walter, Geb- (see Kerr et al., 2011, for a description of participants’ self- hardt, Stark, & Vaitl, 2010). The authors interpret this decrease reports). Questionnaire studies have also documented changes as a diminished involvement in the habitual mode of self- in individuals’ self-concept following mindfulness meditation reference during meditation practice. Meditators show greater practice. Participants’ self-reports of internal and external resting-state functional connectivity within the default mode aspects of self-representation (assessed with the Tennessee network than do nonmeditators (Jang et al., 2010). Another Self Concept Scale; Roid & Fitts, 1988) showed highly sig- study of experienced meditators also reports increased func- nificant changes on almost all of the subscales after comple- tional connectivity between posterior cingulate cortex and tion of a 7-day mindfulness retreat (Emavardhana & Tori, dorsal ACC and dorso-lateral PFC both during rest and during 1997). Changes can be summarized as a more positive self- mindfulness meditation among experienced meditators com- representation, more self-esteem, and higher acceptance of pared with novices, suggesting increased conflict monitoring oneself. A cross-sectional study (Haimerl & Valentine, 2001) and cognitive control over the function of default mode net- that examined the Self-Concept scales of the Temperament work after significant meditation training (Brewer et al., and Character Inventory (Cloninger, Svrakic, & Przybeck, 2011). Although still limited in scope, these studies suggest 1993) of Buddhist meditators with varying levels of medita- that default-mode network activity and connectivity might be tion experience found that increased meditation experience affected in some way by mindfulness meditation practice. was associated with positive development on each of the three Probably the most insightful neuroimaging study to address scales. More experienced meditators showed self-concept the neural correlates of a change in perspective on the self is by styles that are typically associated with less pathological Farb et al. (2007), which investigated brain activity during symptoms. Although these studies do not describe the drastic two forms of self-reference in participants who completed a change in sense of self that highly experienced meditators mindfulness-based stress reduction training; these participants have reported following deep states of meditation, they sug- were compared with another group of individuals who had not gest that some beneficial changes in the perspective on the self yet undergone training; the forms of self-reference included a can happen resulting from mindfulness meditation practice. narrative focus (evaluative monitoring of enduring traits) and an Neuroscientific findings: Functional neuroimaging. Neuroim- experiential focus (momentary first-person experience). Indi- aging studies of mindfulness meditation have demonstrated viduals with the mindfulness training showed larger reductions that brain structures that support self-referential processing in the medial PFC during the experiential (compared with the are structurally and functionally impacted by mindfulness narrative) focus, along with increased engagement of the right meditation. Self-referential processing robustly activates cor- lateral PFC, the right insula, secondary somatosensory cortex, tical midline structures of the brain (Northoff et al., 2006), and inferior parietal lobule. Functional connectivity analyses

Downloaded from pps.sagepub.com at The University of Iowa Libraries on May 24, 2012 How Does Mindfulness Meditation Work? 549 revealed an uncoupling of the right insula and medial PFC, and between each component might seem artificial. We want to there was increased connectivity of the right insula with dorso- illustrate the interaction of the components with an example: lateral PFC regions in the experiential focus after the mindful- During mindfulness meditation, the meditator’s goal is to ness training. The authors interpret these findings as representing maintain attention to current internal and external experiences a shift in self-referential processing, namely, as a shift “toward with a nonjudgmental stance, manifesting acceptance, curios- more lateral prefrontal regions supporting a more self-detached ity, and openness. When an emotional reaction gets triggered and objective analysis of interoceptive (insula) and exterocep- by thoughts, sensations, memories, or external stimuli (e.g., a tive (somatosensory cortex) sensory events, rather than their memory of a frightening event), the executive attention system affective or subjective self-referential value [which is repre- (Section 1) detects the conflict to the task goal of maintaining sented by medial PFC activation]” (Farb et al., 2007, p. 319). a mindful state. Heightened body awareness (Section 2) helps Furthermore, given the higher activity of regions supporting to detect physiological aspects of the feelings present (e.g., body and internal awareness, these data also suggest that body tension, rapid heartbeat, short shallow breath), and the increased body awareness might be closely related to changes in provided information about the internal reaction to the stimu- the perspective on the self, consistent with Buddhist philosophy. lus is a prerequisite for accurate identification of the triggered Greater internal awareness might replace the previous, narrative emotional response (i.e., fear). Emotion regulation processes form of self-reference. (Section 3) then become engaged, in order to relate to the Neuroscientific findings: Structural neuroimaging. In the recent experience differently rather than with a habitual reaction (i.e., longitudinal structural study mentioned above (Hölzel et al., simply noticing the fear as opposed to engaging in avoidance 2011), the posterior cingulate cortex, the temporo-parietal mechanisms). The first two mechanisms (sustained attention junction, and the hippocampus showed increased gray matter [1] to body awareness [2]) lead to a situation of exposure, and concentration following mindfulness-based stress reduction. the third mechanism (regulating for nonreactivity) facilitates Given the relevance of these brain structures for the experi- response prevention, leading to extinction and reconsolidation ence of the self, it seems possible that the structural changes (3). Rather than being stuck in the habitual reactions to the might be associated with changes in the perspective on the external and internal environment, the meditator can experi- self. It is interesting to note that the hippocampus, temporo- ence the transitory nature of all related perceptions, emotions, parietal junction, posterior cingulate cortex, and parts of the or cognitions in each moment of experience. The awareness of medial prefrontal cortex form a brain network (Vincent et al., the transitory nature of the self and one’s momentary experi- 2006) that supports diverse forms of projecting the self onto ence leads to a change in the perspective on the self (4), where another perspective (Buckner & Carroll, 2007), including self-referential processing (i.e., the narrative of the relevance remembering the past, thinking about the future (Schacter, of the stimulus for oneself) becomes diminished, while first- Addis, & Buckner, 2007), and conceiving the viewpoint of person experiencing becomes enhanced. The entire process others, also referred to as a theory of mind (Saxe & Kanwisher, represents enhanced self-regulation, which—according to 2003). These abilities have been suggested to share a common Karoly (1993)—is defined as a process that enables individu- set of processes, by which autobiographical information is als to guide their goal-directed activities by modulation of used adaptively to enable the perception of alternative per- thought, affect, behavior, or attention via deliberate or auto- spectives (Buckner & Carroll, 2007). Structural changes in mated use of specific mechanisms. this brain network (involved in the projection of the self onto As illustrated by the example, the described components another perspective) may be associated with the perceptual mutually facilitate each other. Attention regulation is espe- shift in the internal representation of the self following mind- cially important and, as the basis of all meditation tech- fulness practice. niques, appears to be a prerequisite for the other mechanisms The change in the perspective on the self is precisely to take place. Focused attention on internal events is neces- described in the Buddhist literature but has yet to be rigorously sary in order for practitioners to gain an increased awareness tested in empirical research. The findings reviewed here exem- of bodily sensations with the resultant ability to recognize plify early steps in the process of changing the perspective on the emergence of emotions. The ability to keep attention the self but are still far from addressing the experiences focused on conditioned stimuli is also a prerequisite for the described by highly trained meditation practitioners. Research successful extinction of conditioned responses. Enhanced studies that are currently under way will help elucidate this body awareness might be very closely related to the changes process further and will give us more insight into the underly- in the perspective on the self and might replace a narrative ing neuroscientific mechanisms. form of self-reference. The change in perspective on the self may result in reappraisal of situations in specific ways, which might provide motivation for further development of atten- Integration tion regulation and body awareness. As the components The above described components (see Table 2 for a summariz- mutually facilitate each other, the occurring process could be ing overview) are presumably highly interrelated. In fact, they understood as an upward spiral process (cf. Garland et al., might interact so closely with one another that a distinction 2011).

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Self-Compassion practices vary in the degree to which they foster its increase. Some types of practices are pursued with the primary goal of The concept of self-compassion is closely related to mindful- cultivating (self-) compassion (Germer, 2009; Salzberg, 1995), ness. According to the definition proposed by Neff (2003a), while others strongly emphasize self-compassion within the self-compassion entails three components: self-kindness context of traditional mindfulness meditation (Brach, 2003). (being kind and understanding toward oneself in instances of In mindfulness-based stress reduction, even though it is not perceived inadequacy or suffering rather than being harshly the declared primary goal of the program, self-compassion is self-critical), common humanity (perceiving one’s experi- implicitly and explicitly interwoven into meditation instruc- ences as part of the larger human experience rather than seeing tions, exemplified by reminders included in focused attention them as separating and isolating), and “mindfulness” (in this meditation: “whenever you notice that the mind has wandered context defined as “holding one’s painful thoughts and feel- off, bring it back with gentleness and kindness.” The gentle ings in balanced awareness rather than over-identifying with yoga stretches are practiced with an emphasis on “exploring them” (Neff, 2003a, p. 223). According to Neff’s definition what feels good for oneself and one’s body in this moment.” (2003a), mindfulness thus constitutes one of the three compo- Whenever participants encounter physical pain or emotional nents of self-compassion. suffering, they are encouraged to “take care of themselves.” In line with these practices, self-report studies show that self-compassion scores increase over an 8-week mindfulness- The relationship between self-compassion and based stress reduction course (Birnie et al., 2010; Shapiro, mindfulness Astin, Bishop, & Cordova, 2005; Shapiro, Brown, & In their current conceptualization and operationalization Biegel, 2007; but also see P. D. Abercrombie, Zamora, & within contemporary research contexts (Baer et al., 2006; Korn, 2007). Neff, 2003a), mindfulness and self-compassion are highly cor- related. The total score of the Five Facet Mindfulness Ques- tionnaire and the total score of the Self-Compassion Scale Self-compassion within the theoretical have been found to be correlated (r = .69) in a sample of non- framework proposed here meditators (Hollis-Walker & Colosimo, 2011). Assumptions Within the framework of mechanisms proposed in this study, have been put forth about the nature of their relationship, and self-compassion is presumably most related to emotion regu- it has been suggested that mindfulness is required in order for lation as well as to the change in perspective on the self. The self-compassion to develop because the former enables one to generation of feelings of kindness toward oneself in instances clearly see mental and emotional phenomena as they arise of perceived inadequacy or suffering (self-kindness) is an act (Neff, 2003b). In line with this assumption, changes in mind- of emotion regulation. When cultivating self-compassion, fulness have been found to predict changes in self-compassion seeing one’s difficult experiences as part of the larger human (Birnie, Speca, & Carlson, 2010). It has also been suggested experience rather than seeing them as separating and isolat- that self-compassion partially mediates the relationship ing (common humanity) might initially require reappraisal. between mindfulness and well-being (Hollis-Walker & Colo- This reframing might ultimately result in a change in the per- simo, 2011). Furthermore, the cultivation of self-compassion spective on the self, where one relates to oneself in a less has been suggested to explain much of the success of identified manner. However, there is currently only a very mindfulness-based interventions. Kuyken et al. (2010) found small empirical basis for the explanation of the mechanisms that the positive effects of a mindfulness-based cognitive ther- of self-compassion, and it is possible that unique aspects of apy intervention on depressive symptoms were mediated by self-compassion are not addressed within the suggested the enhancement of self-compassion across treatment. In components. patients with anxious distress, scores on the Self-Compassion We are unaware of any published data on the neural corre- Scale have been found to correlate more strongly with symp- lates of self-compassion in the context of mindfulness training tom severity and quality of life than scores on the Mindful or on the neural basis of self-compassion (but see Lutz, Attention Awareness Scale (Van Dam, Sheppard, Forsyth, & Brefczynski-Lewis, et al., 2008, for altruistic compassion). Earleywine, 2011). However, given the strong interrelatedness However, with the availability of the Self-Compassion Scale of both constructs, it might be difficult to tease their effects (Neff, 2003a), there has been a drastic increase in the investi- and relationship apart. gation of self-compassion in the context of mindfulness-based interventions in the last few years, and a considerable body of literature documents the improvement of self-compassion The cultivation of self-compassion in with mindfulness meditation practice. The question about the meditation practice exact nature of the relationship between both constructs and Meditation is typically practiced with an intention—implicit their interconnection from an empirical and neuroscientific or explicit—to cultivate self-compassion as well as compas- perspective will have to be revisited once more research is sion toward other beings. Different types of meditation available.

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Further Considerations In the progression of meditation expertise, the different mechanisms might play different roles. For example, it is pos- We have suggested here that mindfulness meditation practice sible that an improvement in attention regulation evolves first comprises a process of enhanced self-regulation that can and helps facilitate other processes. Conversely, the change in be differentiated into distinct but interrelated components, perspective on the self might develop rather late, following the namely, attention regulation, body awareness, emotion regula- establishment of increased body awareness and improved tion (reappraisal and extinction), and the change in perspective emotion regulation. Beyond the mechanisms formulated here, on the self. Previous work has often focused on one of these it is possible that increased experience in mindfulness practice components, neglecting the others and attempting to describe facilitates the flexible access to the different components. Pos- the beneficial effects of mindfulness-based interventions sibly the greatest effect of mindfulness practice for adaptive solely through one of the mechanisms (Brown & Ryan, 2003; functioning in daily life might be found in this behavioral Carmody, 2009). Other work has suggested an array of distinct flexibility. components, but these components were not related to one Future clinical psychological research should establish another (Baer, 2003; Brown et al., 2007). Our work in estab- what roles the different components play for different psycho- lishing relations between identified components of mindful- logical disorders. Disorders that manifest as the dysfunction of ness meditation practice is a step toward a more complex a certain component could especially benefit from the cultiva- framework. Such a framework describing a comprehensive tion of that particular component. For example, strengthening process and simultaneously considering the role of subcompo- attention regulation might be most beneficial for patients suf- nents will help advance the field in several ways. First, when fering from attention deficit disorders, while borderline per- conducting basic mindfulness meditation research, differenti- sonality disorder patients, people in addiction recovery, or ating between distinct components will facilitate a more patients with alexithymia might benefit much from increased detailed understanding of the process and stimulate multifac- internal awareness. Likewise, patients with mood disorders, eted research questions. Second, a detailed understanding of anxiety disorders, borderline personality disorder, or aggres- the different components and their relevance for clinical disor- sion might benefit from improved emotion regulation. Change ders will be conducive for the flexible and more targeted in self-perspective might be beneficial for patients with mood application of mindfulness training in psychiatric treatment disorders and might also enhance general sense of well-being and will in turn facilitate the establishment of targeted and as well as overall quality of life in healthy populations. Rather cost-effective programs specifically utilizing components that than solely testing the usefulness of mindfulness-based inter- are most relevant for a specific disorder. Third, a better under- ventions for symptom reduction for these disorders in general, standing of the state and trait effects of mindfulness practice future research should focus more on establishing the mecha- will also be conducive to a better understanding of the func- nisms underlying these beneficial effects. tioning and cultivation of a healthy mind, thereby contributing Different mechanisms might be relevant for different per- to the newly emerging field of positive psychology. sonality types. Individuals likely differ in the extent to which Presumably, the distinct components differ in their rele- they are attracted to the practice of each of these components, vance for types of mindfulness-based meditation practices, and they might differ in the extent to which they can benefit levels of meditation expertise, specific psychological disor- from each of the described mechanisms. For example, it has ders, personality types, and specific situations. In the follow- been found that a self-compassion intervention for smoking ing section, we will suggest connections between the described reduction was particularly beneficial for individuals high in components and these variables. The suggested connections self-criticism and low in readiness to change (Kelly, Zuroff, mostly lack empirical support thus far and are mainly meant to Foa, & Gilbert, 2010). In the same way, individual differences stimulate further research questions. should be taken into account in meditation research. Future Various types of mindfulness practice may place different studies should try to identify traitlike predictors as well as bio- emphasis on the described components. For example, during the logical markers for (a) attraction to specific kinds of practice practice of breath awareness or the body scan (Hart, 1987), the and (b) benefits from particular aspects of such practices. components of attention regulation and body awareness might Aside from differing in relevance between different disor- be most involved. Observing one’s emotions in emotionally ders, personality types, levels of expertise, and types of prac- challenging situations (Kabat-Zinn, 1990) may involve body tices, these mechanisms will have distinct relevance for awareness and extinction. Open awareness practice might different contextual situations. Depending on the kind of situ- mostly involve the change in perspective on self, whereas lov- ation to which a practitioner is being exposed, one of the ing kindness and compassion meditation (Salzberg, 1995) might mechanisms might move into the foreground, while others rely on emotion regulation and the change in perspective on the become less relevant. self. Investigating which components are involved in mindful- Mindfulness as a state, trait, and clinical intervention has ness meditation and which are potentially strengthened by these been extensively researched over the last two decades; how- different types of practice can help individuals in selecting ever, knowledge of the underlying mechanisms of mindfulness which they would like to specifically cultivate. is still in its infancy. Future work should identify additional

Downloaded from pps.sagepub.com at The University of Iowa Libraries on May 24, 2012 552 Hölzel et al. components of mindfulness and establish to what extent the Anand, A., Li, Y., Wang, Y., Wu, J., Gao, S., Bukhari, L., . . . Lowe, components described in this article are truly distinct mecha- M.J. (2005). Activity and connectivity of brain mood regulating nisms or how they can be integrated into fewer components. circuit in depression: A functional magnetic resonance study. Bio- We believe that it will be necessary both to further differenti- logical Psychiatry, 57, 1079–1088. ate each component and to further integrate them into a com- Ancin, I., Santos, J.L., Tijeira, C., Sanchez-Morla, E.M., Bescós, prehensive model. This future empirical work is critical in M.J., Argudo, I., . . . Cabranes-Díaz, J.A. (2010). Sustained atten- order to optimally apply mindfulness in the clinical domain tion as a potential endophenotype for bipolar disorder. Acta Psy- and to advance techniques that aim at cultivating a healthy chiatrica Scandinavica, 122, 235–245. mind and increased well-being. Anderson, N.D., Lau, M.A., Segal, Z.V., & Bishop, S.R. (2007). Mindfulness-based stress reduction and attentional control. Clini- Acknowledgments cal Psychology & Psychotherapy, 14, 449–463. The authors would like to thank Elizabeth Kathleen Avis, Narayan Arzy, S., Thut, G., Mohr, C., Michel, C.M., & Blanke, O. (2006). Brach, Patricia Pop, and Erik Tobiason for their helpful comments on Neural basis of embodiment: Distinct contributions of temporo- this article. parietal junction and extrastriate body area. Journal of Neurosci- ence, 26, 8074–8081. Declaration of Conflicting Interests Asada, H., Fukuda, Y., Tsunoda, S., Yamaguchi, M., & Tonoike, M. The authors declared that they had no conflicts of interest with (1999). Frontal midline theta rhythms reflect alternative activa- respect to their authorship or the publication of this article. tion of prefrontal cortex and anterior cingulate cortex in humans. Neuroscience Letters, 274, 29–32. Funding Austin, J.H. (2006). Zen-brain reflections. Cambridge, MA: MIT Britta K. Hölzel was supported by a Marie Curie International Press. Outgoing Fellowship within the Seventh European Community Baer, R.A. (2003). Mindfulness training as a clinical intervention: A Framework Programme. conceptual and empirical review. Clinical Psychology: Science and Practice, 10, 125–143. Notes Baer, R.A., Smith, G.T., & Allen, K.B. (2004). Assessment of mind- 1. Telomeres are protective DNA sequences at the ends of chromo- fulness by self-report. Assessment, 11, 191–206. somes that ensure genomic stability during cellular replication. Baer, R.A., Smith, G.T., Hopkins, J., Krietemeyer, J., & Toney, L. Telomerase is the cellular enzyme responsible for telomere length (2006). 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